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Priority Actions to Improve Provenance Decision-Making Forum Priority Actions to Improve Provenance Decision-Making MARTIN F. BREED, PETER A. HARRISON, ARMIN BISCHOFF, PAULA DURRUTY, NICK J. C. GELLIE, EMILY K. GONZALES, KAYRI HAVENS, MARION KARMANN, FRANCIS F. KILKENNY, SIEGFRIED L. KRAUSS, ANDREW J. LOWE, PEDRO MARQUES, PAUL G. NEVILL, PATI L. VITT, AND ANNA BUCHAROVA Selecting the geographic origin—the provenance—of seed is a key decision in restoration. The last decade has seen a vigorous debate on whether to use local or nonlocal seed. The use of local seed has been the preferred approach because it is expected to maintain local adaptation and avoid deleterious population effects (e.g., maladaptation and outbreeding depression). However, the impacts of habitat fragmentation and climate change on plant populations have driven the debate on whether the local-is-best standard needs changing. This debate has largely been theoretical in nature, which hampers provenance decision-making. Here, we detail cross-sector priority actions to improve provenance decision-making, including embedding provenance trials into restoration projects; developing dynamic, evidence-based provenance policies; and establishing stronger research–practitioner collaborations to facilitate the adoption of research outcomes. We discuss how to tackle these priority actions in order to help satisfy the restoration sector’s requirement for appropriately provenanced seed. Keywords: assisted migration, ecological restoration, local adaptation, restoration genetics he restoration sector’s demand for seed is Williams et al. 2014, Havens et al. 2015, Prober et al. 2015, Tenormous and is rapidly increasing with the growth Breed et al. 2016b, Christmas et al. 2016b). in the global restoration effort (Verdone and Seidl 2017). Consequently, several alternative provenancing strate- Choosing the geographic origin—the provenance (see the gies have been proposed to supplement local provenances glossary in box 1 for definitions)—of seed used in resto- with nonlocal provenances. These strategies generally fall ration is an early and fundamental decision faced by res- into two categories: those that attempt to increase the adap- toration practitioners (Miller et al. 2017). The traditional tive potential of plants at a restoration site by increasing practice has been to source seed local to the restoration the genetic diversity of the seed mix (e.g., relaxed local, site—local provenancing—because it is expected to maintain composite, and admixture provenancing) and those that the evolutionary history of plant populations (e.g., local recommend matching provenances with anticipated future adaptation), minimize maladaptation, and limit deleterious environmental conditions of a restoration site (e.g., predic- genetic effects (e.g., outbreeding depression). tive and climate-adjusted provenancing). Although there Global change has sparked a broad and robust debate on are theoretical underpinnings to these alternative strate- whether nonlocal seed should be used in restoration and, if so, gies, experimental field-testing in restoration contexts is under what circumstances (Hamilton 2001, Wilkinson 2001, limited. Broadhurst et al. 2008, Mijnsbrugge et al. 2010, Byrne et al. In this article, we identify several knowledge gaps and 2011, Breed et al. 2013, Williams et al. 2014, Havens et al. practical barriers that need priority action by researchers, 2015, Prober et al. 2015, Breed et al. 2016b, Christmas et al. policymakers, and practitioners to improve provenance 2016b, Bucharova 2017). There are two central tenets to this decision-making from an international perspective. We argument: habitat fragmentation and climate change. Habitat discuss these priority actions in light of the global restora- fragmentation generally increases inbreeding and reduces tion challenge of restoring degraded terrestrial landscapes effective population sizes of plant populations, which can from grassland to forest ecosystems, but we acknowledge have an impact on local seed quality by reducing fitness and that the success of a provenancing strategy will be context adaptive potential (Vranckx et al. 2011, Breed et al. 2015). dependent (e.g., landscape heterogeneity and age; plant Because climate is a strong and common agent of selection species biology, demography, and evolutionary history; and in plants (Davis and Shaw 2001, Petit et al. 2008), it has been management priorities, scale, and resources). By highlight- argued that provenance selection should match values of cli- ing these priority actions, we hope to improve support for matic variables predicted into the future (Breed et al. 2013, restoration, offer guidance for policy development, bridge BioScience 68: 510–516. © The Author(s) 2018. Published by Oxford University Press on behalf of the American Institute of Biological Sciences. All rights reserved. For Permissions, please e-mail: [email protected]. doi:10.1093/biosci/biy050 Advance Access publication 7 June 2018 510 BioScience • July 2018 / Vol. 68 No. 7 https://academic.oup.com/bioscience Forum Box 1. Glossary of terms. Admixture provenancing: Supplementing local provenances with seed collections across a species’ natural distribution. Climate-adjusted provenancing: Supplementing local provenances with targeted nonlocal provenances collected along a climate gradient in line with climate-change projections. Composite provenancing: Supplementing local provenances with seed from multiple nonlocal provenances within gene-flow dis- tances to mimic natural gene flow among populations. Cultivar: The selective breeding of a genotype for favorable attributes such as fast growth or high-salinity (or other extreme environ- ment) tolerance. Habitat suitability model: A model of the statistical relationship between species occurrences and environment that conceptually represents the predicted spatial abiotic and biotic limits of a species suitable habitat. These models are also referred to as species distri- bution models or ecological niche models and draw on ecological niche theory. Heterosis: The increased fitness of the hybrid product between two divergent provenances or species compared with parental genotypes. Local adaptation: Superior fitness of a local population over other provenances when grown in its home environment. Local provenance: Seed from either within the restoration site (strict local provenance) or within close geographic proximity to the restoration site (relaxed local provenance). Defining a “local” provenance is a difficult task and is rarely articulated well in papers that discuss this issue, which hinders progress in the provenancing debate. Adopting a genetically informed definition of a local provenance is probably the most useful approach, and examples include spatial delineation of provenances based on significant genetic differen- tiation in neutral and/or adaptive genetic markers or functional traits. However, such an approach is perhaps the least attainable for practitioners because of the costs, time, and skills required. Local provenancing: Only using a local provenance. Outbreeding depression: The decreased fitness of the hybrid product between two divergent provenances due to disruption of locally adapted gene complexes. Provenance: The geographic location of a seed source that is used to describe the genetic material from that location. Provenancing: In an ecological restoration context, refers to a seed-sourcing strategy, with a focus on the geographic location of seed source(s). Phenotypic plasticity: The nongenetic capability of one genotype to produce a variety of different phenotypes. Predictive provenancing: Deriving a matched seed source with predicted future home-site conditions; optimally should be based on provenance trials. Restoration: The process of reducing the damage and/or degradation of an ecosystem by resetting the trajectory of the ecological community toward that of a reference state. This definition should potentially be broadened to capture restoration of sites that do not have a reference (e.g., pollinator habitat in agricultural matrices or biodiverse urban green spaces) and to also include sites that have been degraded to such an extent that restoration to a reference state is likely an unachievable goal (e.g., some postmining sites and novel ecosystems). Seed zones: Discrete geographic regions that have similar environments. Transfer within zones should result in few detrimental effects on mean population fitness. Optimally should be based on progeny testing of multiple provenances. knowledge gaps from science into practice, and ultimately Improving provenance strategy choice. Local provenancing has improve restoration outcomes now and in the future. been the preferred approach for many years. However, a strict interpretation of local provenancing may halt restora- Strategic goals and practitioner priorities tion projects if local seed supply is insufficient (Wilkinson We identify four gaps in provenance decision-making that 2001, Broadhurst et al. 2008). Local provenancing should are commonly faced by both policymakers and practitio- not be so restrictive that it stops projects, even if this means ners. These include (1) the lack of guidance when depart- that nonlocal provenances are introduced. In cases in ing from a strict local provenancing strategy, (2) the need which broader provenances are used or a change in practice for greater evidence-based restoration practices, (3) the is implemented, documentation and ongoing monitor- potential risks associated with using nonlocal
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